Journal of Neuroeducation

Revista de Neuroeducación – Revista de Neuroeducació

Experiences & Electrophysiological (EEG) Correlates

Perspectives
of Reward Effects on Early Sensory
Perception in Humans
Neuroscience, Signal Processing and Psychophysics

Kundan Lal Verma1*

1
Jawahar Lal Nehru Post Graduate College, Deen Dayal Upadhyaya Gorakhpur University,
Gorakhpur, Uttar Pradesh, India; [email protected]

Abstract
Existing research suggests that the reward system and sensory perception net-
works operate in concert and that activation in one can influence the other, but the
dynamics of these influences remain poorly understood. There is general agree-
*
Correspondence ment that there is an interaction between bottom-up and top-down signals in per-
KundanLalVerma
[email protected]
ception and attentional processing. Although it’s not entirely clear which stages of
perception are influenced by cognition, it has been assumed that cognitive input
Citation
Verma KL. Electrophysiological influences later categorization stages of visual processing, and that earlier stages
(EEG) Correlates of Reward Effects are involved only in the pure bottom-up extraction of basic features of sensory
on Early Sensory Perception
in Humans. JONED. Journal of signals. Several recent experiments have challenged this idea by showing that top-
Neuroeducation. 2024; 4(2): down modulation by cognition can extend to early visual stages of perception. Re-
121-127. doi: 10.1344/joned.
v4i1.43569 cent electrophysiological studies have begun to investigate the neural correlates
Reception date: 26/08/2023
of the interaction between attention/reward, perception and cognitive control in
Acceptance date: 04/12/2023 humans. We propose that the selection of the value of our choices and actions
Publication date: 15/02/2024
from multiple alternatives may lead to the suppression of the sensory representa-
Conflict of interest tions of unselected, low-value stimuli, while the selected, high-value stimuli are
The author declare no conflicts of
interest. enhanced. The present project has been proposed to investigate the dynamics
of this selective process by tracking the effects of different reward categories on
Editor
Laia Lluch Molins (Universitat de attention and early sensory perception using behavioural and electrophysiological
Barcelona, España)
(EEG) techniques. Methods from neuroscience, signal processing, psychophysics
Reviewers and EEG tools will be used in the project.
Eric Roig
Olena Klimenko
Keywords: Perception; brain reward system; EEG; sensory; cognition
Copyright
© Kundan Lal Verma, 2024
Resum
This work is licensed under a
Creative Commons Attribution- Les investigacions existents suggereixen que els sistemes de recompensa i les
NonCommercial 4.0 International xarxes de percepció sensorial funcionen conjuntament i que l’activació d’una pot
License.
influir en l’altra; no obstant això, la dinàmica d’aquestes influències encara no es
coneix bé. Hi ha un acord general sobre l’existència d’una interacció entre els se-

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Verma: Electrophysiological (EEG) Correlates of Reward
Effects on Early Sensory Perception in Humans
Experiences & Perspectives

nyals de baix a dalt i de dalt a baix en la percepció i el processament atencional.

Tot i que no està totalment clar quines etapes de la percepció són influenciades
per la cognició, s’ha assumit que l’entrada cognitiva influeix en les etapes de ca-
tegorització posteriors del processament visual, i que les etapes anteriors estan
involucrades únicament en l’extracció pura de baix a dalt de les característiques
bàsiques dels senyals sensorials. Diversos experiments recents han desafiat
aquesta idea mostrant que la modulació de dalt a baix per la cognició pot esten-
dre’s a les etapes visuals primerenques de la percepció. Estudis electrofisiolò-
gics recents han començat a explorar els correlats neurals de la interacció entre
atenció/recompensa, percepció i control cognitiu en humans. Postulem que la
selecció del valor de les nostres eleccions i accions entre múltiples alternatives
pot causar la supressió de les representacions sensorials dels estímuls no se-
leccionats de baix valor, mentre que els estímuls seleccionats de gran valor són
potenciats. El present projecte s’ha proposat per estudiar la dinàmica d’aquest
procés selectiu on es rastregen els efectes de diverses categories de recompen-
ses en l’atenció i la percepció sensorial primerenca utilitzant tècniques compor-
tamentals i electrofisiològiques (EEG). Es faran servir mètodes de neurociència,
processament de senyals, psicofísica i eines d’EEG en el projecte.

Paraules clau: Percepció; sistema de recompensa cerebral; EEG; sensorial; cog-

nició

Resumen
Investigaciones existentes sugieren que los sistemas de recompensa y de per-
cepción sensorial funcionan en conjunto y que la activación en uno puede influir
en el otro; sin embargo, la dinámica de estas influencias sigue siendo poco com-
prendida. Existe un consenso general sobre la existencia de una interacción entre
señales ascendentes (bottom-up) y descendentes (top-down) en la percepción y
el procesamiento atencional. Aunque no está completamente claro en qué eta-
pas de la percepción influye la cognición, se ha asumido que la entrada cognitiva
influye en las etapas posteriores de categorización del procesamiento visual, y
que las etapas iniciales están involucradas únicamente en la extracción pura-
mente ascendente de características básicas de las señales sensoriales. Varios
experimentos recientes han desafiado esta idea al demostrar que la modulación
descendente por la cognición puede extenderse a las primeras etapas visuales
de la percepción. Estudios electrofisiológicos recientes han comenzado a inves-
tigar los correlatos neuronales de la interacción entre atención/recompensa, per-
cepción y control cognitivo en humanos. Planteamos que la selección del valor
de nuestras elecciones y acciones entre múltiples alternativas puede causar la
supresión de las representaciones sensoriales de estímulos no seleccionados y
de bajo valor, mientras que los estímulos seleccionados y de alto valor se realzan.
Este proyecto ha sido propuesto para estudiar la dinámica de este proceso selec-
tivo donde se rastrean los efectos de varias categorías de recompensa en la aten-
ción y la percepción sensorial temprana utilizando técnicas comportamentales
y electrofisiológicas (EEG). En el proyecto se utilizarán métodos de neurociencia,
procesamiento de señales, psicofísica y herramientas de EEG.

Palabras clave: Percepción; sistema de recompensa cerebral; EEG; sensorial; cog-

nición

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Verma: Electrophysiological (EEG) Correlates of Reward
Effects on Early Sensory Perception in Humans
Experiences & Perspectives

Introduction reward value affects early sensory perception in hu-

mans. Previous experimental and theoretical work
Every moment, we regularly experience sensory stim- has predicted that reward-based choices modulate
uli from our surroundings, from the feel of clothes on responses in task-relevant perceptual and cognitive
our skin, to the change of a traffic light from green regions. This sensory modulation provides the sys-
to red, to the trilling of our mobile phone. These sen- tem with the ability to enhance the representation
sory stimuli are often also associated with a certain of high reward stimuli relative to low reward stimuli,
hedonic value, derived from our past experience thereby supporting optimal adaptive behaviour in the
of the positive or negative reward associated with face of changing stimulus-reward associations27, 30.
them20. The reward associations of sensory stimuli How might sensory processing be informed about
are in constant motion, constantly changing as we the reward value of different stimuli? Bottom-up and
unconsciously adapt to the many sensory changes top-down processes are used as effective solutions
that occur in our environment within a single mo- to the limited processing resources of the brain’s
ment1, 3. We are constantly adapting our perception executive system. Bottom-up processing of infor-
to the environment: deciding which rewarding stimuli mation is primarily informed by sensory input. In
are significant and planning our responses to these top-down processing, however, the brain allocates
individual stimuli7. its resources based on behavioural goals and prior
Neuroanatomical studies have shown that the knowledge 2, 13, 32. Previous research has shown that
brain’s central reward pathway sends information top-down processes robustly influence bottom-up
to and receives information from many other brain processing, but the dynamics of these interactions
regions5, 9, 11. The reticular activating system (RAS) di- during reward-based decisions are poorly under-
rects arousal and attention to sensory inputs from stood. Here, we set out to investigate how top-down
our environment. Limbic regions such as the sep- information about the value of different reward cate-
tum, amygdala and thalamus provide input to the re- gories affects bottom-up sensory perception.
ward pathway regarding emotional and motivational The primary goal of this research is to measure
parameters10. The interaction of the reward pathway the effect of different rewards, including food, water,
with the basal ganglia and cerebellum modifies music, visual and auditory tasks, etc., on human per-
the brain’s motor function. Neurotransmitters are ception. The second aim is to define the brain pro-
tracked and controlled by the hypothalamus. This cessing channels that are most frequently engaged
command centre then transmits signals in the form in different cases of reward estimation tasks. Our
of neurotransmitters to different areas of the brain38. main hypothesis is that different reward categories
Electrophysiological methods (such as single cell activate a common reward network, while each spe-
recordings) are widely used to elucidate how differ- cific reward type selectively modulates its respective
ent reward systems influence sensory perception sensory brain area. The transmission of reward value
and action planning. Non-invasive variants of these from the common to the specific areas occurs via
methods, together with advanced data analysis tech- mid-level areas that represent both reward and sen-
niques39, allow proper investigation of brain function sory information. Crucially, we find that this transmis-
in humans. Electroencephalogram (EEG) is an ex- sion can occur by modulating response magnitude
ample of these methods, allowing the measurement as well as by inducing oscillations in a specific fre-
of neural activity time-locked to the presentation of quency band. Using different types of reward allows
stimuli and providing high temporal resolution and us to identify the most commonly used ‘code’ that
dynamics (on millisecond timescales)11, 12, 22. EEG re- different brain areas use to transmit reward informa-
cordings could also be performed simultaneously tion across the brain.
with measurements of hemodynamic changes in the The old study by Gilchrist and Nesberg in 1952
brain, as in fMRI, which provides high spatial resolu- showed that hungry subjects overestimated the
tion (spatial resolution in millimeters)4, 25. These meth- brightness of pictures of food compared to other
ods are used in various psychological, behavioural pictures. Later, in 1957, Bruner’s conceptual basis of
and clinical neuroscience experiments19, 29, 37. the New Looks framework suggested that perception
The motivation of this review is to investigate how is influenced by top-down factors related to individ-

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Verma: Electrophysiological (EEG) Correlates of Reward
Effects on Early Sensory Perception in Humans
Experiences & Perspectives

ual expectations and needs. In 1997, Breiter’sHuman • Data Acquisition

studies of rewarding stimuli suggested that intensely Recording
pleasurable emotions are mediated by neural activi- EEG signal recorded by wired or wireless recording
ty in neural systems through motivational, emotion- systems on multiple channels as required by proto-
al (limbic) and arousal processes. In 2001, Anne J. col or event.
Blood experimented with music rewards and found
a link between physiological signals and the inten- Preprocessing
sity of the music 6.Another line of evidence comes This step includes normalisation, digitisation and fil-
from combined behavioural and EEG studies that tering. Normalisation is a very important step; it only
have investigated the dynamics of sensory process- minimises the scaling of the signal without changing
ing. A study by Sophie Molholm in 2002 suggested its basic characteristics. The raw EEG data is filtered
that reaction time changes according to audio-vi- for noise induced by power line interference, EMG
sual neural interactions23.Recent studies show that and EOG etc33, 34, 36. The task of filtering is carried out
electroencephalography (EEG) provides a method to by implementing various filters such as band pass
highlight the temporal characteristics of synesthetic filters, low pass filters, high pass filters and adaptive
experience in order to evaluate early sensory pro- filters depending on the quality of data and analysis
cessing5, 16, 24. A recent experiment by Porbadnigk in requirements. Digitisation of data is done to convert
2013 used a machine learning approach on EEG data the analogue data into digital data (however, it is not
to investigate brain states in auditory perception 31. required in case of already digitised recording devic-
Similarly, a number of studies have investigated the es).Analogue signals are continuous and smooth.
EEG correlates of perception in the visual modality Digital signals represent discrete points in time and
(for a review see: Norcia et al.)26. A separate line of their values are quantities with a fixed resolution rath-
research has sought to investigate the neural cor- er than those of continuous signals. Digitised signals
relates of reward processing in humans. Hagerty are convenient for data analysis purposes36.
used both fMRI and EEG to investigate the reward
system14.Krigolson et al. 2013 showed that learning • Feature Extraction
of reward associations in a gambling task is explic- Extracting the relevant data from the digitised data
itly reflected in changes in evoked EEG responses to that can be used for proper classification is an im-
novel reward cues over time17, 21.Importantly, recent portant step 8. The extracted feature contains the
studies have shown that during perceptual tasks, useful information, therefore the accuracy of the
sensory stimuli associated with reward elicit chang- feature extraction affects the accuracy of the clas-
es in evoked potentials or the oscillation patterns of sification. Feature extraction can be done in both
EEG data15, 18 compared to unrewarded stimuli. These time and frequency domain. Some examples of time
findings suggest that changes in the synchronous domain analysis are common spatial pattern (CSP)
activity of distributed neuronal cell assemblies may evaluation, auto-regressive parameter estimation,
underlie the effects of reward on perception. Our pro- basic probability and statistical analysis 34. Some
posed idea of correlating EEG parameters with early examples of frequency domain analysis are Fourier
sensory perceptual functions across different reward spectral analysis and power spectral density estima-
categories is inspired by this idea and is unique in tion. Data reduction can be achieved in the feature
its conceptualisation, falling within the latest trends extraction process by selecting only the relevant data
in neuroscience research (for a review of the impor- for classification.
tance of elucidating reward effects on early sensory
perception see 27, 28). • Classification
Once the correct features have been extracted, they
Materials and methods need to be identified as a particular occurrence or
class using classification algorithms35, 36.
The EEG signal analysis method is used in the case
of reward effects on early sensory perception.EEG • Source localization of EEG signals
signal processing involves the following steps The method of source localisation of EEG signals

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Verma: Electrophysiological (EEG) Correlates of Reward
Effects on Early Sensory Perception in Humans
Experiences & Perspectives

is used to localise brain activity; it provides useful tory). Sensory processing of each reward catego-
information about physiological, psychological and ry will be selectively modulated when a perceptual
functional abnormalities of the brain. This problem judgement is required about another exemplar of
is called the EEG inverse problem36. the same category (but not the other categories). As
a result of this research, we will be able to identify
Data collection analysis and evaluation a general brain reward processing system across
60-100 healthy subjects of different age groups, male different reward categories. This is a crucial step in
and female, left and right handed, will be recruited the development of strategies to enhance sensory
to participate in studies on the effects of reward on performance and cognitive control in humans, for
perception through different reward systems. All example where sensory processing is impaired (e.g.
subjects will be required to sign the informed con- poor vision) or pathologically hypersensitive (e.g. in
sent form to participate in these studies. They may addiction).
be asked to complete a questionnaire to establish
their profile. Prior to the start of the experiment, the Discussion
subjects will be given a training session to familiarise
them with the do’s and don’ts of the specific research The environment is rich in different types of informa-
protocol. tion. Sensory organs such as the eyes, ears, tongue,
The normal baseline EEG parameters recorded skin and nose provide sensory information such as
will be compared with the EEG parameters recorded light, sound, taste, touch and smell. This information
during or after reward-based decisions, so that the not only informs the brain about the sensory proper-
effect of reward learning can be assessed. Reward ties of environmental stimuli, but also signals their
based decisions involve the presentation of differ- associated rewards. In fact, these two types of in-
ent reward categories, with the subject indicating formation can influence each other, and behavioural
which stimulus they choose (by pressing a button). studies have repeatedly demonstrated these interac-
We compare EEG correlates (in the time or frequency tions.
domain) across different reward categories to identi-
fy those that are common across reward classes and Conclusions
those that are specific to each class.
The general brain reward processing system will
Results help us to understand the effects of different ear-
ly sensory perceptions in human subjects. We will
We expect that oscillations in a specific frequency also test which of these EEG signals correlate with
band carry reward information throughout the brain, behavioural performance measured by reaction time
regardless of the reward category (visual or audi- and choice probability.

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Experiences & Perspectives

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Experiences & Perspectives

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